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l&A
THE WOLVES OF YELLOWSTONE
Natural Resources Report
Number 14
U.S. Department of the Interior
National Park Service
Digitized by the Internet Archive
in 2014
https://archive.org/details/wolvesofyellowst243weav
THE WOLVES OF YELLOWSTONE
John Weaver
Department of Wildlife Science
Utah State University , Logan
and
Environmental Research Institute
Moose, Wyoming
U.S. DEPARTMENT OF THE INTERIOR, NATIONAL PARK SERVICE
Natural Resources Report Number 14 • 1978
For sale by the Superintendent of Documents. U.S. Government Printing Office,
WMhington, D.C. 20402
Stock No. 024-006-00712-1
ABSTRACT
Historical records and intensive field surveys 1975-77
provided information on the population history, ecology, and
current status of wolves (Canis lupus) in Yellowstone National
Park and vicinity. Wolves occurred in unknown but seemingly
low densities during the latter 1800s in several areas of
Yellowstone where they were controlled periodically until 1926.
Populations apparently began increasing about 1912, primarily
in the northeast, and may have reached nonequilibrium levels
of 30-40 animals (postwhelping) . Intensive control 1914-26
removed at least 136 wolves, including about 80 pups. During
this period Yellowstone wolves characteristically lived in
packs of 3-16 members, some of which followed the ungulates
in their seasonal migrations. Litters averaging 7.8 were born
in late March and April, primarily in the north central sector
of the park. Limited evidence suggests that elk (Cervus elaphus)
were important food for wolves during all seasons. Wolves either
survived the control era or moved in shortly thereafter for
singles, pairs, and a pack of four were reported the following
decade. Resident wolf packs, however, were eliminated from
Yellowstone National Park by the 1940s. Large canids have
been sighted intermittently to the present, but their identity
has not been established. Singles and pairs comprised 89% of
116 "probable" reports over the past 50 years. Speculation
about factors limiting the Yellowstone wolf population considers
its relative geographic isolation from viable wolf populations
and possible genetic problems (including wolf-coyote hybridiza-
tion) associated with prolonged minimal population status. A
transplant of wolves from British Columbia or Alberta, or perhaps
Minnesota, is recommended to restore a viable population of
this native predator to Yellowstone National Park.
Cover drawing by Carol Snow
As the Nation 's principal conservation agency, the Department
of the Interior has responsibility for most of our nationally
owned public lands and natural resources. This includes
fostering the wisest use of our land and water resources ,
protecting our fish and wildlife, preserving the environmental
and cultural values of our national parks and historical
places, and providing for the enjoyment of life through out-
door recreation. The Department assesses our energy and
mineral resources and works to assure that their development
is in the best interests of all our people. The Department
also has a major responsibility for American Indian reserva-
tion communities and for people who live in Island Territories
under U.S. administration.
THE WOLVES OF YELLOWSTONE
INTRODUCTION
Wolves historically occupied a wide range
of habitats throughout much of North America
north of the 20th parallel in southern
Mexico (Goldman 1944) , but their geographi-
cal range in the contiguous United States
today has been reduced by nearly 991 (Mech
19 "1). The Northern Rocky Mountain Wolf
(NRMW) (C. I. ivremotus) , one of 23 sub-
species recognized by Goldman, once roamed
the backbone of the continent from southern
Idaho and Wyoming to southeastern British
Columbia and southern Alberta (Fig. 1).
Wolves throughout this area, including
Yellowstone National Park, were reduced
drastically by the 1930s by government and
private control. In 1973 the Secretary of
the Interior placed the NRMW on the
Endangered Species List.
The primary purpose of administration of
natural areas by the National Park Service
is to preserve natural environments and
native plant and animal life while provid-
ing for enjoyment by visitors in ways which
maintain natural conditions (USDI National
Park Service 1968) . Mission-oriented re-
search involves determining the complete-
ness of park ecosystems and developing
management procedures to prevent or compen-
sate for departures caused by human actions
(Cole 1969a) . Lack of ecological complete-
ness, for example, might stem from unnatu-
ral reduction or elimination of predator
populations .
In recent years personnel and visitors
in Yellowstone National Park have reported
sightings of large canids (Cole 1971) . No
intensive field research, however, had been
conducted specifically on wolves there.
Mech (1971) stated:
For the wolves reported from Yellow-
stone Park, an immediate and concerted
program is necessary. Intensive
efforts should be made to determine
the extent of populations . . .
both in terms of numbers and area
occupied. Special attention should
be given to determine whether breed-
ing and successful reproduction are
taking place.
In August 1975, I was contracted as an
independent research biologist by the
National Park Service to survey the status
of wolves in Yellowstone National Park.
Objectives of the study were to compile the
historical information on wolves in Yellow-
stone and to determine their present dis-
tribution, abundance, and reproductive
success .
1
1 . Cants lupus tundrarum
2. C. I. fambasileus
3. C.l. dees
4. C.l. occi den talis
5. C. I. hudsonicus
6. C. I. arctos
7. C. /. orion
8. C. I. Ubradorius
9. C. I. btotkuctu
10. C. /. I j coon
11. C. /. nub 'dus
12. C. /. irremotus
13. C . I. eolumbianus
14. C. /. ligoni
15. C. /. fuscus
16. C. /. crassoion
17. C. /. youngi
18. C. /. mogoUonensii
19. C. /. monstrabilu
20. C. /.
21. C. /. bernarii
22. C. /. mackenzii
23. C. /. marsningi
FIGURE 1. North American distribution of subspecies of Canie Zupua (from Goldman 1944).
2 National Park Service
STUDY AREA
A 1?.1S1 km2 (7,481 mile2) area, encom-
passing Yellowstone National Park and a 16-
24 km strip around its perimeter in north-
western Wyoming and adjacent parts of
Montana and Idaho (Fig. 2) , was selected
for the wolf survey. Much of the area is
designated or de facto wilderness, and de-
velopments such as roads, buildings, and
campgrounds occupy less than 1% of the
park.
Quaternary volcanic deposits which have
undergone three glaciations cover most of
the area (Keefer 1972). Elevations range
from about 1,500 m to over 3,400 m, but
forested rhyolite plateaus at 2,100-2,600
m are extensive.
Winters are usually long and cold while
summers are short and cool (Dirks 1976;
Houston 1976) . Most of the annual precip-
itation of 34.5-96.5 cm falls as snow. In
general, temperatures are lower and precip-
itation higher in the central and southern
parts of the area.
About 19% of the terrestrial area of
Yellowstone is forested, with lodgepole
pine v04i) and subalpine fir-Engelmann
spruce (6%) predominating. Despain (1973)
and Houston (1976) have described the vege-
tation.
Distribution and estimated abundance of
bison (Bison bison) (Meagher 1973) , elk
(Cole 1969b; Craighead et al. 1972; Houston
1974), moose {Aloes aloes), mule deer
(Odocoileus hemionus) , pronghorn antelope
(Antilocapra americana) , and bighorn sheep
(Ovis canadensis) (Barmore in prep.) have
been reported (Table 1) Ecology of tue
cyote {Canis latrans) (Murie 1940) ana
grizzly bear {Ursua arctos) (Craighead et
al. 1974; Mealey 1975; Cole 1976; Knight
et al. 1977) has been presented. Houston
(1973) commented upon tne status of mountain
lions (Felis conoolor) and wolverines (Julc
aula) in the park.
METHODS
Information on the history and ecology
of wolves in Yellowstone National Park up
to the 1930s was obtained from journals,
Army scout diaries, Army station records
(extracted by M. Meagher) , and from monthly
and annual reports of the superintendent,
in the Yellowstone National Park Research-
Reference Library.
Population trends since that time to the
present were assessed from nearly 500 re-
ports of wolf-like animals and/or sign.
These reports included replies to a ques-
tionnaire mailed to 89 big-game outfitters
operating on the Gallatin, Shoshone, and
Teton National Forests adjacent to the park.
A point system was devised for evaluating
and categorizing the observations (Table 2).
The principal criteria included experience
and reliability of the observer, details of
the observation, and description of the
animal and/or sign which would distinguish
it in external appearance from other canids.
Reports were categorized as "probable" or
"possible," depending upon the number of
points received. A "positive" category
was reserved for instances where an animal
was trapped or killed and verified as a
wolf.
This point system was designed to be
conservative. For example, an observation
of a large gray canid at short range by a
person familiar with Western coyotes re-
ceived a "possible" evaluation. Reports
citing a distinctive color, howl, or track
rated "probable" if the observer was
Natural Resources Report No. 14
3
FIGURE 2. Location of the Yellowstone wolf study area. Broken lines arbitrarily distin-
guish five geographical sections to facilitate discussion of reported wolf observations.
Shaded circles represent ungulate baits and/or canid scent and time-lapse cameras,
February- March 1977.
National Park Service
TABLE 1. Seasonal population estimates of ungulates in
Yellowstone National Park; 1977.
Species Summer Winter
Elk 20,000-25,000 10,000-13,000
Mule deer 2,000- 4,000 500- 2,000
Moose 1,500 1,000
Bison 1,100- 1,200 1,100- 1,200
Bighorn sheep • 600 600
Pronghorn antelope 150 150
qualified. All reports were classified
independently of any others.
This system remains subjective, however,
and with such sources, all reports are
questionable to a degree. Some observa-
tions, especially those from the 1930s and
1940s, may well have been of wolves but
were classified "possible" for lack of de-
tails. Some "probable" sightings may have
been of large coyotes. Nonetheless, the
point system provides consistency for
evaluating these observations. All reports
were coded on computer-compatible sheets
and filed at park headquarters.
I spent 12 months in the field, August-
October 1975 and August 1976-April 1977,
searching intensively for wolves and/or
sign. Since the wolves' reproductive suc-
cess was an important question, I designed
the field study to cover periods of breed-
ing, denning, and rendezvous activity. My
field assistants and I traveled approxi-
mately 2,700 km on foot, skis, and snow-
shoes while inspecting government-maintained
trails, game trails, ridges, and stream
courses for wolves, tracks, and scats.
Tape-recorded and human- imitated wolf
howls were broadcast approximately 1,400
times both day and night from elevated
spots (Joslin 1967). A parabolic microphone
was available for recording any responses.
Baits of road-killed ungulates and canid
scent were placed at seven locations in and
near the park (Fig. 2), February-March 197",
and were monitored with time- lapse movie
cameras (Diem et al. 1973). Some cameras
were preset to expose a single- frame pic-
ture at 1.5-minute intervals; others, at
8 -minute intervals.
Thirty hours in flight time were spent
searching specifically for wolves and/or
sign. In addition, approximately 1,800
hours have been logged by other park re-
search biologists since 1964 during wild-
life distribution and censusing flights.
Most of the intensive ground search con-
centrated on the northeast and southeast
portions of the study area (Fig. 2), where-
as flights were made over most of the park.
THE YELLOWSTONE WOLF
The wolves of Yellowstone- -C. I.
irremotuB - -probably intergraded with colum-
bianuB to the north, nubilus to the east,
and youngi to the south (Goldman 1944) .
Specimens from northwestern Wyoming were
considered by Goldman to be "somewhat
Natural Resources Report No. 14
5
TAE^r 2. Criteria and point system for categorizing wclf
observations .a
Criteria
Points
Observer
experience with Western coyotes
experience with wolves
Observation
distance
<100 m
10C-4C0 m
>400 m
length of observation time
>10 seconds
optical aid such as binoculars
Description of animal and/or sign
body description
large body size
large and blocky head, short ears,
and relatively short muzzle
relatively large, long legs
color
solid white or black
h ~ wl
distinctively different from coyote
track
>10 cm (4.0 in) long including
toenails (must include evidence
to rule out domestic dog)
- 7
3
u
2
10
10
10
aReports scoring £16 points qualified as "probable";
those <16 points, "possible." See text for discussion
of system.
intermediate" between irremotua and youngi.
Goldman (1944:404) pointed out that gray
wolves "... are all very similar in the
more essential features and are believed to
intergrade through the vast range of the
species on the North American mainland."
Indeed, taxonomists today, with multivari-
ate statistical techniques, might reduce
the number of wolf subspecies or perhaps
eliminate them altogether (see Nowak 1973;
Jolicoeur 1975; Skeel and Carbyn 1977) .
Although there are reasons to question the
6
National Park Service
validity of the subspecific groupings, I
use Northern Rocky Mountain Wolf (XRMQ for
convenient reference to this geographical
population. Wolf taxonomy and its implica-
tions for management will be discussed
later.
The NRMK is medium- to large-sized for
the species. An adult male from Red Lodge,
Montana, measured 1,870 mm (61.4 inches)
total length, while two adult females from
Soda Springs, Idaho, measured 1,929 and
2,046 mm (Goldman 1944). An adult rale
taken in Montana in 1968 weighed 42 kg (92
lb) (Gary Day pers. comm.). For wolves in
Yellowstone, Bailey (1930) stated: "The
male is consistently larger than the female,
weighing well over a hundred pounds."
Goldman (1944) described the winter
pelage of the NRMW:
Upper parts from nape to rump usually
near "light buff" or varying shades
of tray, sparingly overlaid with
black, becoming nearly white cn
riivf- and limbs; short pe_age cr. top
if r.ead light buffy white, the nairs
tipped with black; ears and upper
surface of muzzle light buffy; under
parts in general more cr less soiled
white; tail -above light buffy, thinly
and inconspicuously overlaid with
black, light buffy below to tit,
which is a mixture of buff and slack
all around. Individuals in the black
phase appear to be rare.
Of 136 wolves killed in the park, three
were black and one was white (Fig. 3); two
others observed were black. All others
were gray. It seems reasonable that most
black or white wolves, due to their con-
spicuousness, would be reported. Neither
the reported kills nor the recorded obser-
vations substantiate Skinner's (1927) claim
that up to 40?0 of Yellowstone wolves were
black.
POPULATION HISTORY
Prior to 1914
Wolves were members of Yellowstone's
native fauna. Although few observations
were recorded during the 1800s (Appendix
I), this could reflect either an actual low
density of wolves or simply a lack of rec-
ords. Some early writers used 'Voir* in
reference to both true wolves and coyotes
("small prairie wolf" or "medicine wolf)
{see Haines 1955:129), but reports from
about 1880 on usually distinguished the two
species. Five accounts of gray wolves were
recorded 1869-80.
At least as early as 1877, however,
ungulate carcasses in the park were poisoned
with strychnine by free- lance "wolfers" for
"wolf or wolverine bait" (Supt. .Annual
Rept. 1877). By 1880, Superintendent
Norris stated in his annual report that
". . . the value of their [wolves and
coyotes] hides and their easy slaughter
with strychnine-poisoned carcasses have
nearly led to their extermination."
Sightings of single wolves, pairs, and
groups of three and six were recorded 1881-
1908, primarily in the Geyser Basins (C) ,
Hayden Valley (C) , and Lamar (NE) (Fig. 4).
The Yellowstone wolf population appar-
ently began to increase about 1912, at
least in the northern sector. In a letter
dated 29 July, 1912, Col. L. M. Brett
remarked that "... several were killed
on the Upper Gallatin River but a few miles
outside [the park] . . . last spring"
(Appendix I). Skinner (1927) observed four
wolves in Lamar Valley that same year and
itural Resources Report No. iu
7
ft
TZZ'SPX 3. Three wolves killed near Hellroaring Creek ( NE ) and the den in a rock cave from
which six pups were removed, Yellowstone National Park, April 1916 (from Bailey 1916).
riGURL 4. Location of wolf observations in Y
circles represent sightings of singles or pai_
together.
believed wolves "were coming in faster."
Randall (19661 saw a pack of nine along the
Yellowstone River near Hellroaring in
spring, 1915.
1914-1926
By 1914, wolves had increased noticeably
in northeast Yellowstone Park (Supt. Annual
Rept. 1914). They were considered, though,
"a decided menace to the herds of elk, deer,
mountain sheep, and antelope" (Supt. Annual
Rept. 1915) and concerted efforts to "exter-
minate" (Supt. Monthly Rept. February 1919)
wolves were mounted. Opposition to this
policy was ignored as suggested by the
llowstone National Park prior to 19lu. Open
s; shaded circles, three or more animals
following (Supt. Monthly Rept. May 1922):
"It is evident that the work of controlling
these animals must be vigorously prosecuted
by the most effective means available
whether or not this meets with the approval
of certain game conservationists."
During 1914-26 a minimum of 136 wolves--
• including about 80 pups (59*) —were removed
from dens, trapped, shot, and probably
poisoned within the park (Table 3). This
total is slightly higher than Skinner's
(1927) and Murie's (1940) due to a more
detailed examination of historical sources.
In each of 3 years (1918, 1920, 1922), the
toll exceeded 24 animals (mostly pups) . In
Natural Resources Report No. 14
FIS'JRE 5. Wolf pups trapped at bison carcass near Soda Butte (NE), Yellowstone Park,
October 1926. (Photo by Scott Riley.)
FIGURE 6. Location of observations and numbers of wolves killed (X) in Yellowstone Park
and vicinity, 191^-25. Open circles represent sightings of singles or pairs; shaded
circles, three or more animals together.
10 National Park Service
TABLE 3.
Wolf mortality
in Yellowstone
National Park, 191-
-2C .
Totals"
Year -
Adults
Pups
Monthly Repts. A:
r.r.. r.ept .
191U
7
-
7
-
1915
3
4
7
"several"
1916
2
8
10
12
1917
2
-
2
-
1918
21
iua
35
36
1919
2
2
6
— V,
28
1920
6
20+
26+
1321
3
11
1U
1922
9
16
25
24
1923
1
5
6
8
1924
1925
1926
2
2
Total
56
80
136
114
Report did not distinguish between adults and pups. Nineteen
wolves were killed in April and I have assumed that, as for the
same month in other years, most of these were pups.
b
One den with unreported number of pups was closed up.
CCombined for 1921 and 1922.
^Monthly reports were more detailed and were considered to be
the best source.
1918, 21 adult wolves were killed. The
annual kill fluctuated throughout this
period, indicating either real changes in
wolf numbers or varying effort by the con-
trol personnel, or both. Certain trends,
however, appear evident.
Between 1914 and 1923, 15 (56*) of 27
reports involved three or more wolves to-
gether (Appendix II) . Four occupied and
distinct dens found in 1916 and 1920 sug-
gest at least four different reproductive
units those years. The last den destroyed
by park personnel was in 1923 near Tower
Falls (NE). In the next 3 years, only 3
(21%) of 14 reports mentioned more than two
wolves together. Two pups were trapped near
Soda Butte (NE) in October 1926 (Fig. 5) .
All the reported killing of wolves oc-
curred in the northeast section from near
Mammoth east to Soda Butte and south to
Pelican Valley (Fig. 6), as did many obser-
vations. In addition, one or two wolves
were sighted occasionally in the southeast
section. Hayden Valley and the Geyser
Basins, areas occupied by wolves around the
turn of the century, furnished but one
record of a single animal during 1914-26.
Natural Resources Report No. 14
11
FIGURE 7a. Locations of canid observations in Yellowstone Park and vicinity, 1927-36.
Open circles represent sightings of singles or pairs; shaded circles, three or more
animals together.
1927-1966
The population history of wolf-like
canids in Yellowstone 1927-66 relies upon
reports which have been classified
"probable" and Superintendent Monthly
Reports up to 1936. Within the recognized
subjective nature of the data, this repre-
sents my best interpretation of wolf popu-
lation trends during those years. To
facilitate discussion, I grouped reports
into 10- year periods.
1927-26. In the decade following intense
persecution of wolves in the park, 14
observations of 29 large canids were
tallied. Seven reports involving 17 ani-
mals were classified "probable" (Fig. 7a)
(see Appendix III). A pack of four was
observed up Tower Creek (NE) in 1934
(Arnold 1937), but sightings of singles
or pairs accounted for five of seven
reports.
1937-46. There were 16 reports of 18 large
canids for this period; 8 of these invol-
ving 10 animals were rated "probable"
(Fig. 7b). Three wolves were seen in 1937
on lower Specimen Ridge, a single on Soda
Butte Creek in 1938, and another single
just north of the park in 1942 (all in NE) .
A wolf was observed up Mol Heron Creek (NW)
in 1942 and another up the Gallatin River
the following year. In 1944, single wolves
were reported at Heart Lake (SE) , Elk Park
(NW) , and Crevice (NE) . Sightings of
singles or pairs comprised seven of eight
reports.
12
National Park Service
FIGURE 7b. Locations of canid observations in Yellowstone Park and vicinity, 1937-46.
Open circles represent sightings of singles or pairs; shaded circles, three or more
animals together.
1947-56. Twenty-six reports of 37 canids
were received; six involving 10 animals
were classified "probable" (Fig. 7c). Two
wolves were observed at Soda Butte in 1947,
one near Lost Creek in 1949, and three at
Amethyst Creek in 1952 (all in NE) . A
single, large canid was seen just south of
the park in 1950. Along the Madison River,
a pair of wolves was seen in 1952 and a
single, the following year. Five of these
six "probable" reports mentioned only sin-
gles or pairs.
1957-66. Thirty-two observations of 42
canids were recorded. Fourteen reports
involving 21 animals received a "probable"
rating (Fig. 7d) . Single wolves were
observed in the northeast in 1957, 1958,
1963, and 1965. A large adult canid with
three pups was reported there in 1963.
Singles were seen near Swan Lake (NW) in
1958 and 1965 and a pack of three on the
Gallatin River in 1966. One of two wolves
observed up Mol Heron in 1963 was shot but
not recovered. In 1960 one wolf was re-
ported at Grouse Creek (SE) south of
Yellowstone Lake and another, along the
east boundary near Sunlight Basin (NE) . A
single wolf was seen near Porcupine Hills
(C) in 1963. Sightings of singles and pairs
constituted 12 of 14 reports.
Surmary. Wolf- like canids either survived
the 1914-26 control era in Yellowstone
National Park or moved in shortly there-
after. Murie (1940) believed that "the
last wolves were eliminated in the twenties
although a few have been reported in recent
Natural Resources Report No. 1U
13
FIGURE nz. Locations of canid observations in Yellowstone Park and vicinity, 19U7-56,
Open circles represent sightings of singles or pairs; shaded circles, three or more
animals together.
FIGURE 7d. Locations of canid observations in Yellowstone Park and vicinity, 1957-66.
Open circles represent sightings of singles or pairs; shaded circles, three or more
animals together.
National Park Service
FIGURE 8. Locations of canid observations in Yellowstone Park and vicinity, 1967-April
1977. Open circles represent sightings of singles or pairs; shaded circles, three or
more animals together.
years." Two packs of three to four and
five wolves may still have been present in
the mid- 1930s, but records do not indicate
they persisted. During the next 30 years,
observations of wolf- like canids --mostly
singles and pairs --were reported sporadi-
cally from the northwest and northeast sec-
tions of the study area.
POPULATION STATUS
1967 -April 1977
During the past decade, 401 reports of
531 canids have been received; 81 involving
109 animals were classified Mprobable,,
(Fig. 8). The increase in reports during
this period was due partly to a system
established in 1968 for recording sightings
of wolves (Cole 1971) and to greater aware-
ness of their possible occurrence in the
park.
About 901 of the "probable" observations
came from four areas (Fig. 8). Each year
throughout this period, one or two wolf-
like canids have been seen in the northeast.
In Hayden Valley (C) , a single wolf was
observed in 1969 and 1971. A wolf reported-
ly was shot in Sunlight Basin east of the
park in January 1968, but no remains were
found the following summer. Sightings of
a large canid in that area were recorded in
1968, 1969, 1971, and 1975. In the north-
west section of Yellowstone, one to five
wolves were seen in 1968-70, a pair in 1971,
and singles in 1972 and 1974. None has
been reported from that area since. Sixty
Natural Resources Report No. 14
15
(74°) of the "probable" observations oc-
curred during 1963-71. Singles and pairs
accounted for 91$ of tnese reports over the
entire period.
During approximately 1,800 hours of
flight by park biologists over all sections
of the park since 1964, only one wolf- like
canid has been seen. D. Houston and D.
Stradley observed this dark animal in
Hayden Valley in 1971, but poor light con-
ditions ,-recluded a positive identification.
Durir.^ 12 months I found only two sepa-
rate sets of tracks and heard one series of
howls which may have been wolf. All oc-
curred on the Shoshone National Forest,
Wyoming, within 20 km of each other and
1-22 km east of the park boundary. Tracks
measuring 11.3 cm (4.5 inches) long (in-
cluding nails) by 9.6 cm (3.8 inches) wide
with a 63- cm stride were found in sand near
a stream on 7 September 1975. No evidence
of human activity or domestic dogs was
discovered anvwhere in the general area.
Tape-recorded wolf howls were broadcast the
previous and following nights, but no re-
sponses were heard. Two weeks later, one
canid howled three separate times in re-
sponse to a broadcast wolf howl. This oc-
curred about 18 km from the site of the
tracks. The animal was in dense timber
and never was observed. No measurable
tracks were found as the ground was hard
and dry. The howls were not recorded, but
I am confident the animal was not a coyote.
Subsequently, 34 days were spent afield in
this area (August -September 1976, April
1977). Canid tracks, identical in measure-
ment to the first set, were found 20 April
1977 about 2 km from the site of the howls.
Casts of these tracks are in the park
Museum Collection.
No wolves were photographed by time-
lapse cameras monitoring the ungulate baits
and/or canid scent during February-March
1977, nor were any observed during 30 hours
of flight. Unfortunately, winter cond.nif.ns
were very mild and many wildlife species
remained scattered throughout the study
area.
ECOLOGY
Reproduction
Breading and Denning Dates. The earliest
positive birth date known for wolves in
Yellowstone was 26 March (1916) when pups,
judged less than 1 week old, were taken
(Appendix II) . Pups were removed from dens
in March 1920 and 1921, but the exact date
was not recorded. Skinner (1927) reported
"three lots of pups which were born about
March 1." Pups of unreported age were
taken from other dens 1-8, 16, and 30 April,
and 12 May. For wolves in the Bighorn
Basin of north-central Wyoming, King (1965)
related denning dates of 25 and 29 March
and 19 April. Assuming a 63-day gestation
(Brown 1936; Woolpy 1968), these dates indi-
cate that Yellowstone wolves bred anytime
from January until early March. This breed-
ing season coincides with others reported
from a similar latitude (Mech 1970:117).
Litter Sizes. The size of 10 presumably
complete wolf litters extracted from dens
averaged 7.8 and ranged from 5 to 13 (Fig.
9) . Litters of 11 and 10 were found in
1921 and 1922, respectively, following
several years of persecution. Such large
litters seem characteristic of exploited
wolf populations (Mech 1970) .
Dens. Bailey (1930) stated that wolf dens
in Yellowstone were usually "situated in
caves or hollows among rocks or sometimes,
16
National Park Service
3r
8
10 II
12
13
Number of Pups in Litter
FIGURE 9. Histogram of litter sizes of wolves in Yellowstone Park, 1916-23.
in large burrows on steep hillsides." He
described a den near Hellroaring as com-
posed of four or five large burrows dug
into the open hillside "which, evidently,
had beer, used for several years." When
disturbed by humans, the adults moved the
pups to another den in a natural cave about
a nule ;' i . 6 km) away (see Fig. 3).
Certain physiographic features appear
characteristic of these and other wolf den
sites described in the literature (Mech
1970:120-121; Stephenson 1974). Typically,
dens are located on south or southwest
aspects of moderately steep slopes in well-
drained soils (or rock caves) , at eleva-
tions 2-200 m above the surrounding area,
and usually within 30-200 m of surface
water. All the reported wolf dens in
Yellowstone were located in the north
central part of the park, from Blacktail
Deer Plateau to Specimen Ridge (Fig. 10).
Rendezvous Sites. Murie (1944) used the
term "rendezvous" for specific resting and
gathering areas occupied by wolf packs dur-
ing summer after the natal den has been
abandoned. These are usually small, open
meadows close to wooded cover and surface
water (Joslin 1967; Carbyn 1974). In August
1922, Park Ranger Anderson found an apparent
rendezvous site of wolves in Yellowstone.
This is a section of the park that is
practically inaccessible due to bog
holes, rim rock, down timber, and
jack pines [sic]. The area is the
part of the Mirror Plateau lying
near the head of Timothy, Raven,
Pelican, and Broad Creeks [Appendix II].
It is possible that this area had been used
by wolves for rendezvous in previous years.
Nowlin (1912) reported wolves howling at the
head of Raven Creek on 25 July 1912. Bailey
(1930) found tracks "especially numerous
along Pelican and Raven Creeks where at
least ten or a dozen wolves hunted in one
pack in July and August, 1915."
Natural Resources Report No. 14
17
Pack Size
Wolves characteristically live in packs
of three or more individuals (Mech 1970) .
Although a pack usually functions as an in-
tact unit, members may split off temporar-
ily at any season of the year (Murie 1944;
Burkholder 1959; Jordan et al. 1967). Hence,
cursory observations of wolves may under-
estimate true pack size.
Nonetheless, even rough estimates of
pack si;? may provide insight if, as Rausch
(1967) proposed, pack size reflects popula-
tion density. Between 1902 and 1926, wolf
packs of 3-16 members were reported for 9
different years. Based upon the number of
distinct dens occupied by wolves, three to
four reproductive units were present in
some years (1916-22) in the northeast. By
contrast, between 1927 and 1977, there have
been "probable" reports of three to five
wolf- like canids together in only 8 years.
It is doubtful if more than one group ex-
isted in any one year.
Seasonal Distribution and Movements
Some wolves in Yellowstone apparently
followed the ungulates in their altitudinal
migrations to and from summer and winter
ranges. Bailey (1930) reported that "during
the summers of 1914 and 1915 they [wolves]
. . . were following the elk herds to the
high pastures of Mirror Plateau, returning
with them in winter to the valleys along the
Lamar and Yellowstone Rivers." The Super-
intendent's Monthly Reports during 1918
state:
Towards the end of the month [May]
the wolves seemed to leave the
Specimen Ridge district and have
not been much in evidence since.
They were considerably in evidence
in Slough and Hellroaring Creeks
[November] .
Although some wolves wintering in the Lamar
and Yellowstone valleys moved toward Mirror
Plateau and Pelican Valley during summer,
FIGURE 10. Approximate location of wolf dens in Yellowstone Park, 1916-23.
18
National Park Service
others may have headed north out of the
park.
Food Habits
Kills by wolves and scat contents sug-
gest that elk were important prey both win-
ter and summer for wolves in Yellowstone.
Scout McBride found a cow elk killed on 21
December 1914, between Mammoth and Black-
tail Deer Creek. Between 16 October and 31
January 1916, Scout Black discovered eight
elk killed by wolves (Appendix II) . Skinner
(1927) wrote that during the winter of
1914-15, two or three wolf packs "harried"
the elk on the lower valleys of the park.
Bailey (1930) reported that wolf droppings
in Pelican Valley collected during July
1915 were made up entirely of elk hair.
That same month he also discovered a young
elk in Slough Creek which he believed had
been killed by wolves.
Studies of food habits of wolves in the
Rockey Mountain National Parks of Canada
(Cowan 1947; Carbyn 1974) and in Glacier
National Park in Montana (Singer 1975) pro-
vide an interesting comparison with Yellow-
stone since similar species of prey inhabit
these areas. Cowan (1947) reported that
elk hair occurred in 49% of winter scats
and 42% of summer ones. Mule deer hair
was found in about 15% of scats collected
at both seasons. Most wolf kills found
by personnel were either elk (54%) or mule
deer (23%). Cowan believed the actual
kill of elk may have been higher because
very few scats were collected in areas
where wolves subsisted "almost exclusively
on elk." Carbyn (1974) found elk hair in
46% of 1,190 summer scats and 11% of 265
winter scats. Mule deer hair was detected
more often (66%) in the winter samples
than in the summer (301) . His study area
contained more mule deer than other areas
of Jasper Park where Cowan (1947) worked.
In both Canadian studies, bighorn sheep
and mountain goats (Oreamnos americanus)
were comparatively invulnerable to wolf
predation. In Montana's Glacier Park,
Singer (1975) reported eight white- tailed
deer (Odocoileus virginianus) , three moose,
one elk, one beaver ' (Castor canadensis),
and five snowshoe hares [Lepus americanus)
killed by wolves.
Elk were also important food for Yellow-
stone wolves during the denning season in
late March and April. On 26 March 1916,
scouts Black and Stevenson found "a score
or more of old elk skulls . . . and one
fresh elk head" near a den in Hellroaring.
A freshly killed young elk was discovered
about 0.5 km from another den in the same
area. Scraps of elk meat were in the den
with the pups (Appendix II) . Considering
the abundance of beaver between Hellroaring
and Tower Falls around 1920 (Warren 1926)
and the reported predilection of wolves
for them (Voight et al. 1976), it seems
likely they formed a portion of the wolves'
diet, too. At two wolf dens in Jasper
Park, Cowan (1947) found remains of 12 elk,
2 mule deer, and 2 beaver. Carbyn (1974)
reported occurrence of mule deer in 44%
and elk in 32%- of 312 scats collected at
wolf dens. Later in summer at rendezvous
sites, 55% of 270 scats contained elk,
while 22% had mule deer.
Data on the sex and age of ungulates
killed by wolves in the Rockey Mountains
are limited but suggest that calves or
fawns and individuals 10 years and older
may be most vulnerable. Of nine elk
killed in Yellowstone and reported by
scouts, six were adult cows, two were
Natural Resources Report No. 14
19
calves, and one was unidentified. No kills
of adult bulls were recorded (Appendix II).
In Jasper, Cowan (1917) classified 66 wolf-
killed elk: very young--20, mature--29,
and diseased-senile- -17. Carbyn (1974) be-
lieved that young individuals less than 3
years old of all ungulates were most vulner-
able to wolf predation. Individuals 3-9 '
years old appeared relatively secure.
DISCUSSION
Woives inhabited the Yellowstone area in
unknown densities when the park was estab-
lished in 1872 but were subject to early ex-
ploitation (1870s) and later control (1914-
26) . A noticeable population increase
about 1912 was met by intense year-round
control, especially removal of pups from
dens. Estimates of wolf numbers, based
upon population stability via reproductive
responses to removal of wolves S months and
older (Rausch 1967: Kelsall 1968; eec Mech
1970) , cannot be made from the Yellowstone
data. Nonetheless, certain comments seem
appropriate. Control records (Table 3) and
the presence of up to four reproductive
units (Appendix II) suggest postwhelping
populations of 30-40 wolves around 1920,
primarily in the northeast and southeast
(Fig. 6). Of 103 wolves observed 1914-23,
83% were in packs of 3 or more. What level
the Yellowstone wolf population might have
reached had control been minimal remains
unknown.
After wolf control ceased within the
park in 1926, very few wolves were reported.
Whether these were remnant survivors or
immigrants, or both, is unknown. Wolf
numbers elsewhere in Wyoming and Montana
were reduced drastically by this time, too.
In Wyoming and South Dakota 508 wolves
were killed 1918-23 by government personnel,
with the last one taken in 1940 (G. Rost
pers. comm.). In Montana, government agents
removed 413 wolves during 1918-30 and their
last one in 1945 (N. Miner pers. comm.).
Yellowstone records do not indicate that any
resident wolf packs persisted after the mid-
1930s.
Wolf-like canids have been sighted with-
in the study area intermittently to the
present, with an increase of "probable"
reports 1968-71. Based upon geographical
distribution of the sightings and some
pelage differences, up to 10 of these canids
may have occupied several separate areas
around 1970. Observations reflect human
distribution and variations in the visi-
bility of animals as well as their actual
seasonal ranges. Hence, it is difficult
from these cursory reports to determine
whether these canids were residents or
transients.
The identity of wolf- like canids re-
ported recently in Yellowstone Park has
not been established and would require
skeletal material. Interpretations include:
(1) a remnant population of genetically
pure wolves persisted through occasional,
successful recruitment; (2) wolves in
Canada and Montana immigrated periodically
down to Yellowstone; (3) wolves were re-
leased into the park; and (4) wolf-like
canids occurred through hybridization of
various kinds.
The interpretation that a small popula-
tion of pure wolves survived seems unten-
able without more consistent evidence of
pack activity during the intervening years.
Use of toxicants on surrounding public
and private lands would have made journeys
hazardous for immigrating wolves. In
Glacier National Park, black wolves, more
20
National Park' Service
prevalent in C. I. "colurbianus" (Canadian)
than in "irremotus , " comprised 12% of the
wolf observations (Singer 1975). Near
Three Forks, Montana (100 km northwest of
Yellowstone Park) , a wolf with a very large
skull was killed in 1941, and both Goldman
(1944) and Cowan (1947) believed it a
Canadian emigrant. But' in Yellowstone
from 1966-77, S% of canids sighted were
black, suggesting that few Canadian wolves
immigrated as far south as Yellowstone.
Despite suggestions of sub rosa releases
of wolves (Mech and Rausch 1975), park of-
ficials have denied that any were ever
transplanted to Yellowstone. Although I
found no evidence to the contrary, this
allegation has not been fully investigated.
The possibility of a surreptitious release
of captive wolves by private individuals
cannot be totally discounted, but their
chances for survival would seem slight.
rhe prolonged geographical isolation of
Yellowstone from wolf population centers
suggests the remaining possibility--
hybridi ration. While reproductive isola-
tion between species is usually maintained
by geographic and ethological barriers,
individuals on the periphery of their
species* range may have trouble finding a
conspecific mate. In the absence of ade-
quate breeding stimuli, they may respond
to inadequate signals and hybridize (Mayr
1963). Recent evidence from several areas
of North America suggests that coyotes
may hybridize both with gray wolves
(Kolenosky 1971; Mengel 1971; Kolenosky
and Standfield 1975; Lawrence and Bossert
1975; Hilton 1976) and red wolves (Canis
rufus) (Paradiso and Nowak 1971; Riley
and McBride 1972; Gipson et al. 1974;
Elder and Hayden 1977) . Coyote-dog
crosses are considered much less likely
in Yellowstone due to its remoteness and
the peculiar reproductive timing of such
hybrids (Mengel 1971).
Kolenosky (1971, 19"" pers. comm.) re-
ported that Fl offspring of an Ontario wolf
( $ ) and coyote {(f) phenotypically resem-
ble Eastern coyotes but with massive legs
and large feet. He suspected that if they
were sighted in the wild they would be
identified as "normal" Eastern coyotes.
However, in external appearance, one of the
F2,s is almost identical to an Algonquin-
type wolf {see Kolenosky and Standfield
1975), with larger head, legs, and feet
than other progeny. Some wild canids in
Ontario (Kolenosky and Standfield 1975) and
Maine (Hilton 1977) appear to be mainly
coyotes with some introgression of wolf
genes. In Yellowstone we have no cranial
material with which to trace such an incur-
sion, if it ever occurred. Yet many re-
ports describe animals phenotypically
similar to the F1 canids observed by
Kolenosky. Some of the recorded track mea-
surements (9-11 cm long x 7-9 cm wide) are
between typical coyote and wolf in size.
Whatever the identity of some large
canids in Yellowstone, the sporadic nature
of reports and the high incidence (89* 6f
116 "probable" sightings) of singles and
pairs over the past 50 years do not indi-
cate a viable wolf population in the park.
MANAGEMENT RECOMMENDATIONS
A stated purpose of the National Park
Service is to "conserve, perpetuate, and
portray as a composite whole the indige-
nous . . . terrestrial fauna" (USDI
National Park Service 1968) . A departure
from natural conditions exists in Yellow-
stone National Park because fewer pure
Natural Resources Report No. 14
21
wolves, if any, occur now than in the past.
Control bv humans- -both within and outside
the park--has brought the Yellowstone wolf
to the edge of extinction.
Two options are available for wolf man-
agement in Yellowstone National Park: (1)
do nothing; or (2) attempt to restore a
viable wolf population by introduction.
The former alternative has been employed
since 1927 when wolf control ceased in the
park. Over the next 50 years, a viable
population has not reestablished, and the
wolf niche appears essentially vacant.
Therefore, I recommend restoring this na-
tive predator by introducing wolves to
Yellowstone.
In proposing a transplant, one must
consider the suitability of source stock.
The Department of the Interior originally
placed C. I. irremotuB on the Endangered
Species List. However, use of trinomens
has been questioned for many animals
(Wilson and Brown Brown and Wilson
1954; Hagmeier 195S; Chapman and Morgan
1973), including wolves (Mech 1974; R. M.
Nowak pers. com.). Recent multivariate
analyses of wolf skulls (Nowak 1975;
Jolicoeur 1975; Skeel and Carbyn 1977)
have shown few statistically significant
differences between many subspecific group-
ings made by Goldman (1944). Upon consid-
eration of such factors, the U.S. Fish and
Wildlife Service has proposed (Federal
Register, 9 June 1977:29527) deleting
C. 1. irremotus and listing the entire
species (Ccris lupus) as endangered
throughout the 48 contiguous states (ex-
cept Minnesota) .
Perhaps a more important aspect of re-
introduction is finding wolves that would
have the best chance of adapting to the
physiography and prey of Yellowstone.
Wolves trom the mountains of British
Columbia or Alberta would seem suitable,
especially if gene flow from that direction
has occurred. Wolves from Minnesota are
another possible source.
22
National Park Service
REFERENCES
ARNOLD, M. 1937. Yellowstone wolves. Nature Mag. Yellowstone Natl. Park, Wye. August: 111-1
BAILEY, V. 1930. Animal Life of Yellowstone National Park. Charles C. Thomas, Baltimore.
241 p.
BARMORE, W. J. In prep. Population characteristics, distribution, and habitat relationships
of six ungulates in Northern Yellowstone National Park.
BLACK, C. 1916. Scout diary, 1915-16. Yellowstone Natl. Park Arch.
BROWN, C. E. 1936. Rearing wild animals in captivity, and gestation periods. J. Marrmal.
17:10-13.
BROWN, W. L.- and E. 0. WILSON. 1954. The- case against the trinomen. Syst. Zool. 3:174-176.
BURKHOLDER, B . L. 1959. Movements and behavior of a wolf pack in Alaska. J. Wildl. Manage.
23:1-11.
CARBYN, L. N. 1974. Wolf predation and behavioral interactions with elk and other ungulates
in an area of high prey diversity. Canad. Wildl. Serv. Rept., Edmonton. 233 p.
CHAPMAN, J. A. and R. P. Morgan II. 1973. Systematic status of the cottontail complex in
western .Maryland and nearby West Virginia. Wildl. Monogr. No. 36. 54 p.
COLE, G. F. 1969a. Mission-oriented research in the natural areas of the National Park
Service. Res. Note 6. Yellowstone Natl. Park. 6 p.
. 1969b. The elk of Grand Teton and southern Yellowstone national parks. Res. Rept.
GRTE-1. Yellowstone Natl. Park. 192 p.
. 1971. Yellowstone wolves. Res. Note 4. Yellowstone Natl. Park. 6 p.
. 1976. Management involving grizzly and black bears in Yellowstone National Park,
1970-75. Nat. Res. Rept. No. 9. 56 p.
COWAN, I. M. 1947. The timber wolf in the Rocky Mountain national parks of Canada. Can.
J. Res. 25:139-174.
CRAIGHEAD, J. J., G. ATWELL, AND B. W. O'GARA. 1972. Elk migrations in and near Yellowstone
National Park. Wildl. Monogr. No. 29. 48 p.
, J. R. VARNEY, and F. C. CRAIGHEAD, Jr. 1974. A population analysis of the Yellow-
stone grizzly bears. Mont. For. and Conserv. Expt. Sta. Bull. 40. Univ. Montana,
Missoula. 20 p.
DESPAIN, D. 1973. Major vegetation zones of Yellowstone National Park. Yellowstone Natl.
Park Info. Paper No. 10. 4 p.
DIEM, K. L. , L. A. WARD, and J. J. CUPAL. 1973. Cameras as remote sensors of animal
activities. Univ. Wyoming, Laramie. 10 p.
DIRKS, R. A. 1976. Climatological studies of Yellowstone and Grand Teton National Parks,
continuing studies. Dept. Atmospheric .Sciences, Univ. Wyoming.
ELDER, W. H. and C. M. Hayden. 1977. Use of discriminant function in taxonomic determina-
tion of canids from Missouri. J. Marrmal. 58:17-24.
GIPSON, P., J. A. SELANDER, and J. E. DUNN. 1974. The taxonomic status of wild Canis in
Arkansas. Sy3t. Zocl. 23:1-11.
GOLDMAN, E. A. 1944. The Wolves of North America, Part II: Classification of Wolves.
Am. Wildl. Inst., Washington, D. C, p 389-636.
HAGMEIER, E. M. 1958. Inapplicability of the subspecies concept to the North American
marten. Syst. Zool. 7:1-7.
HAGUE, A. 1893. Notebook No. 1. Natl. Arch. (USGS library).
HAINES, A. 1955. Osborne Russell's Journal of a Trapper. Oregon Hist. Soc, Portland. 179
. 1965. The Valley of the Upper Yellowstone. An Exploration of the Headwaters of
the Yellowstone River in the Year 1869. Univ. Oklahoma Press, Norman. 70 p.
HENDERSON, A. B. 1870. Narrative of a prospecting expedition to the East Fork and Clark's
Fork of Yellowstone. Yellowstone Natl. Park Lib. p 96.
HILTON, H. 1976. Physical characteristics, taxonomic status and food habits of the eastern
coyote in Maine. M.S. Thesis, Univ. Maine, Orono. 76 p.
HOUGH, E. 1894. Forest and Stream's Yellowstone Park Expedition of 1894. Newspaper Acct.
Yellowstone Natl. Park Lib.
HOUSTON, D. B. 1973 (Rev.). Cougar and wolverine in Yellowstone National Park. Res. Note
5. Yellowstone Natl. Park. 22 p.
23
1974. The northern Yellowstone elk, Parts I fill: history and demography. Yellow
stone Natl. Park. 1S5 p.
. 1976. The northern Yellowstone elk, Parts III 6 IV: vegetation and habitat rela-
tions. Yellowstone Natl. Park. 444 p.
JOLICOEUR, P. 1975. Sexual dimorphism and geographical distance as factors of skull
variation in the wolf Canie lupus L. Pages 54-61 in M. F. Fox, ed. The Wild Canids.
Van Nostrand Reinhold, New York.
JONES, W. A. 1875. Report on the Reconnaissance of Northwestern Wyoming, Including
Yellowstone National Park, Made in the summer of 1873. Government Printing Office,
Washington, D. C. 331 p.
JORDAN, P. A., P. C. SHELTON, and D. A. ALLEN. 1967. Numbers, turnover, and social struc-
ture of the Isle Royale wolf population. Am. Zool. 7:233-252.
JOSLIN, P. W. B. 1967. Movements and home sites of timber wolves in Algonquin Park. Am.
Zool. 7:279-288.
KEEFER, •*. R. 1972. The Geologic Story of Yellowstone National Park. U.S. Geol. Surv. Bull
1347. 92 p.
KELSALL, J P. 1968. The Migratory Barren-Ground Caribou of Canada. Can. Wildl. Serv.
Monogr. No. 3. Ottawa. 340 p.
KING, C. L. 1965. Reasons for the Decline of Game in the Bighorn Basin of Wyoming.
Vantage Press. New York. 161 p.
KNIGHT, R. R., J. BASILE, K. GREER, S..JUDD, L. OLDENBURG, and L. ROOP. 1977. Annual Rept.
Interagency Grizzly Bear Study Team, 1976. 75 p.
KOLENOSKY, G. B. 1971. Hybridization between a wolf and a coyote. J. Mammal. 52:446-449.
, and R. 0. STANDFIELD. 1975. Morphological and ecological variation among gray
wolves (Cants lupus) of Ontario, Canada. Pages 62-72 in M. F. Fox, ed. The Wild Canids
Van Nostrand Reinhold, New York.
LAWRENCE, B. and W. H. BOSSERT. 1975. Relationships of North American Canis shown by
multiple character analysis of selected poulations. Pages 73-86 in M. F. Fox, ed.
The Wild Canids. Van Nostrand Reinhold, New York.
MAYR, E. 1965. Animal Species and Evolution. Harvard Univ. Press, Cambridge. 797 p.
McBRIDE, J. 1914. Scout diary. Yellowstone Natl. Park Arch.
MEAGHER, M. M. 1973. The Bison of Yellowstone National Park. NPS Sci. Monogr. Ser. No. 1.
161 p.
MEALEY, S. P. 1975. Tiu. natural food habits of free ranging grizzly bears in Yellowstone
National Park, 1973-74. MVS-." Thesis, Montana State Univ., Bozeman. 158 p.
MECH, L. D. 1970. The Wolf: The Ecology and Behavior of an Endangered Species. Natural
History Press, New York. 389 p.
. 1971. Where the wolves are and how they stand. Nat. Hist. 80(4):26-29.
. 1974. Canis lupus. Mammalian Species No. 37, p 1-6. Am. Soc. Mammalogists, New
York.
, and R. A. Rausch. 1975. The status of the wolf in the United States— 1973. In:
Wolves. IUCN Suppl. Paper No. 43. Morges, Switzerland. 145 p.
MENGEL, R. M. 1971. The study of dog-coyote hybrids and implications concerning hybridiza-
tion in Canis. J. Mammal. 52:316-336.
MURIE, A. 1940. Ecology of the Coyote in the Yellowstone. Fauna of the Natl. Parks, No. 4
Government Printing Office, Washington, D.C. 206 p.
. 1944. The Wolves of Mount KcKinley. Fauna of the Natl. Parks, No. 5. Government
Printing Office, Washington, D. C. 238 p.
NOWAK, R. M. 1973. North American quaternary Canis. Ph.D. Thesis, Univ. Kansas, Lawrence.
380 p.
NOWLIN, D. C. 1912. Report on wild buffalo and elk, Yellowstone Park, July 22-30. 5 p.
In: File no. 21 6 22, Yellowstone Natl. Park Arch.
PARADIS0, J. L. and R. M. NOWAK. 1971. A report on the taxonomic status and distribution
of the red wolf. USDI Spec. Sci. Rept. Wildl. 145:1-36.
RANDALL, L. W. 1966. Footprints Along the Yellowstone. The Naylor Co., San Antonio. 186
RAUSCH, R. A. 1967. Some aspects of the population ecology of wolves, Alaska. Am. Zool.
7:253-265.
RILEY, G. A. and R. T. McBRIDE. 1972. A survey of the red wolf Canis rufus. USDI Spec.
Sci. Rept. Wildl. 162:1-15.
SINGER, F. J. 1975. Status and history of timber wolves in Glacier National Park, Montana.
Syrop. Wolf Behavior and Ecology, Wilmonton, N. C. 17 p.
SKEEL. M. A. and L. N. CARBYN. 1977. The morphometric relationship of gray wolves
(Canis lupus) in national parks of central Canada. Can. J. Zool. 55:737-747.
National Park Service
SKINNER, M. P. 1927. The predatory and fur-bearing animals of the Yellowstone National
"ark. Roosevelt Vildl. Bull. 4(2) : 163-281 .
STEPHENSON, R. 0. 1974. Characteristics of wolf den sites. Final Rept. Job 14.6 R,
Alaska Dept. Fish 6 Game, Juneau. 27 p.
STEVENSON , D. 1916. Scout diary, 1915-16. Yellowstone Natl. Park Arch.
USDI NATIONAL PARK SERVICE. 1968. Administrative Policies for Natural Areas of the
National Park System. Government Printing Office, Washington, D. C.
VOIGHT, D. R., G. B. Kolenosky, D. H. Pimlott. 1976. Changes in summer foods of wolves
central Ontario. J. Vildl. Manage. 40(4): 663-668.
WALWORTH, L. 1971. Interview with E. R. Hall, Univ. Kansas, Lawrence. Yellowstone Natl
Park Biol. Off. files.
WARREN, E. R. 1926. A study of beaver in the Yancey Region of Yellowstone National Park
Roosevelt Vildl. Ann. 1:1-191.
WILSON, E. 0., and W. L. BROWN. 1953. The subspecies concept and its taxonomic applica-
tion. Syst. Zool. 2:97-111.
W00LPY, J. H. 1968. The social organization of wolves. Nat. Hist. 77:46-55.
ACKNOWLEDGMENTS
This study was funded by the National Park Service,
National Audubon Society, Boone 6 Crockett Club, Wyoming
Environmental Institute, N. J. Bellegie, M.D., and J. F.
Turner through the NPS Cooperative Park Studies Unit, Utah
State University, Logan, and the Environmental Research
Institute, Moose, Wyoming. Drs. Thadis Box (USU) , Douglas
Houston (NPS) , and Frank Craighead (ERI) assisted with
administrative matters. Yellowstone Park personnel were
very helpful and supportive, especially Douglas Houston,
Mary Meagher, Glen Cole, Vicky Kurtz, Richard Knight, and
the Ranger Division. Mary Meagher generously permitted
use of her unpublished notes from archival material. All
the material in the appendixes, except monthly and annual
reports of the Superintendent and published references,
were extracted by Dr. Meagher. The Gallatin and Shoshone
National Forests provided logistical support. Kirk Knudsen,
Jan Peterson, and Marguerite Deimel ably assisted in the
field. Dave and Roger Stradley, Gallatin Flying Service,
provided excellent flight service. Susan Sindt prepared
the figures and Carol Snow drew the cover for this publication.
Drs. I. M. Cowan, F. C. Craighead, Jr.., D. B. Houston, F. F.
Knowlton, and M. M. Meagher read the manuscript and made
helpful suggestions. To each of you, my sincere gratitude.
Natural Resources Report No. 14
APPENDIX I
Summary of Wolf Reports Prior to 1914, Yellowstone National Park.
Source
Date
ReDort
Haines (1955)
Haines (1965)
Henderson (1870)
Jones (1875)
Supt. Annual
Report (1877)
Supt. Annual
Report (1880)
Supt. Annual
Report (1881)
Hague (1893)
Hough (1894)
Soda Butte
Station Record
Fountain Station
Record
Lake Station
Record
Fountain Station
Record
1836
1869
16 September
1870
1873
6 August
1877
1880
1881
1886-87
1894
March
1899
1 June
3 June
13 June
29 June
1901
13 November
19 November
1902
20 September
1902
1 November
Trapper Osbom Russell heard a
hovl near the outlet of Yellow-
stone Lake (could be either wolf
or coyote.
Howl heard at Cache Creek.
Group (wolves) seen at junction
of Cache Creek with Lamar River.
Doleful haul of a large wolf —
near Pelican Meadows.
Ungulate carcasses poisoned with
strychnine for wolves.
Hides of wolves taken in late
fall. The large ferocious gray
or buffalo wolf, the sneaking,
snarling coyote, and a species
apparently between the two of
a dark-brown or black color,
were once exceedingly numerous
in all portions of the Park,
but the value of their hides
and their easy slaughter with
strychnine-poisoned carcasses
of animals have nearly led to
their extermination.
Howl heard in Hayden Valley
Wolf in Upper Geyser Basin.
Billy Hofex saw wolves in
Hayden Valley.
Wolf seen near Slough Creek.
Wolf seen near Slough Creek.
One wolf seen on northeast side
of Slough Creek just above
Buffalo Creek.
Three wolves seen on southwest
side Slough Creek.
Wolf tracks seen at Lower Geyser
Basin.
Wolf tracks seen at Bear Park.
Six wolves seen between Lake and
Mud Geyser.
One wolf seen at Goose Lake.
27
Appendix I
Wolf Reports Prior to 1914
Source
Date
Report
Fountain Station
Record
Soda Butte
Station Record
Sylvan Pass
Station Record
Upper Basin
Station Record
Letter from W. B.
Sheppard to
Col. L. M. Brett,
dated 29 Jan. 1912
Letter from
Col. L. M. Brett
to Win. J. Homaday
Nowlin (1912)
Walworth (1971)
Supt . Annual
Report (1912)
M. P. Skinner
(1927)
Randall (1966)
3 November
18 November
1904
21 April
1907
3 September
4 September
1908
10 August
1911
Late August
1912
29 July
1912
25 July
1912
December
1912
1912
1913
Spring
Two wolves seen at north end of
Mesa Road on the Gibbon River
One wolf seen at 8-mile post
between Fountain and Riverside.
Two wolves between Fort Yellow-
stone and Yancey's.
Two wolves seen between Sylvan
and Lake.
One wolf seen between Lake and
Sylvan.
One wolf seen between Upper
Basin and Excelsior.
Gray wolves, of which latter
I sawy two, and considerable
sign . . .
McBride has been in the Park
for many years, and is not
convinced that there have ever
been any gray wolves here.
Statements have been made
that they have been seen, but
none have ever been killed or
captured inside of the Park
though several were killed on
the Upper Gallatin River but
a few miles outside in the
state of Montana, last spring.
Wolves howling at head of
Raven Creek.
Tracks of 3-4 wolves seen at
Buffalo Ranch (Lamar).
It is claimed that gray wolves
have been heard and that their
tracks have been seen in the
Park, but up to this time none
have ever been killed, and there
is no absolute proof that they
exist within the limits of the
reservation, though they have
been taken not many mv les out-
side on the cattle ranges in
Montana.
In 1912, I saw four [wolves]
near Lamar Valley. After that,
signs of their presence increased
and I believed they were coming
in faster.
Randall saw pack of nine (wolves)
along Yellowstone River trail
near He 11 roaring Creek.
National Park Service
APPENDIX II
Summary of Wolf Reports 1914-26, Yellowstone National Park.
Source
Date
Report
James McBride
(1914)
Lake Station
Record
Skinner (1927)
Letter of trans-
mittal from F. T.
Arnold, Captain
12th Cavalry, to
Sec. of Interior
McBride (1914)
Letter of trans-
mittal from F. T.
Arnold, Captain
12th Calvary, to
Sec. of Interior
Supt. Annual
Report (1914)
Skinner (1927)
Vernon Bailey
(1930)
1914
29 January
1914
25 April
1914
7 September
1914
3 December
1914
21 December
1914
31 December
1914
1914-15
1914 and
1915, Summer
Tracks of three wolves between
Mammoth and 10 miles west.
One wolf seen between Lake and
Pelican Creek.
. . . When I found an extra-
ordinarily bold pack of eleven
big fellows [wolves) in the
Pelican Valley.
Four wolves "killed by Ranger
Henry Anderson on Slough Creek.
Wolves have become rather
numerous along the north line
of the Park during the past two
or three years, and have been
seen frequently, but this is
the first instance where anyone
has been able to capture them
or get close enough to shoot
them.
One cow elk killed by wolves
between Mammoth and Blacktail
Deer Creek.
Three more wolves nave been
killed in the Park during the
month making a total of seven
killed, and there are indications
that they are present in con-
siderable numbers and are
destroying much game.
Gray wolves have made their
appearance in the Park in
considerable numbers, having
been seen traveling in packs of
ten or less. While efforts have
been made to kill them, thus far
none have been taken inside of
the Park although a few have
been killed just outside, along
the northern border . . . efforts
will be made to kill them.
That winter, two or three packs
harried the elk on the lower,
open valley 8 of the Park ....
They began to increase about
1914, soon numbered about
sixty ....
During the summers of 1914 and
1915 they [wolves] . . . were
following the elk herds to the
high pastures of Mirror Plateau,
Natural Resources Report No. 14
Appendix II
Wolf Reports 1914-26
Source
Date
Report
Vernon Bailey
(1930) - Cont.
1915
July
Bailey (1930)
1915
November
1915
returning with them in winter to
the valleys along the Lamar and
Yellows tone Rivers. In the
summer of 1915, Mr. Frazier, at
the Buffalo Ranch [Lamar}, told
me that wolves had been very
troublesome during the preceed-
ing winter and had killed many
elk. During June of that yeaTi
Mr. Frazier killed two half'
grown wolf pups and caught two
more, which were kept chained
up at the ranch. During July
and August, 2315, I found where
a family of wolves had killed
and eaten a young elk in Slough
Creek Valley and found wolf
tracks along Slough Creek and
Lamar Valleys up to the mouth of
Mist Creek, also along Pelican
Creek, and later a few tracks
on Fox Creek at the southern
edge of the Park. Tracks were
especially numerous along
Pelican and Raven Creek where
at least ten or a dozen wolves
hunted in one pack.
On this same trip I found big
wolves common, feeding their
young on elk, and probably also
on buffaloes, as they were right
in the midst of the buffalo ranges.
This probably accounted for the
slow rate of increases of the
herd, for after the wolves were
trapped out of this section the
following winter by Donald
Stevenson, the herd began to
make rapid increase.
Donald Stevenson counted nine
separate tracks, where a band of
wolves had crossed a sandbar on
Pelican Creek, but at that time
they were leaving that section
of the Park and following the elk
herds to lower levels.
On Pelican Creek, along the trails
which they [wolves] were constantly
using, their droppings were made
up entirely of elk hair, and a
scarcity of elk calves was very
noticeable among the herds in
that section.
National Park Service
Appendix II
Wolf Reports 1914-26
Source
Date
Rercrr
Bailey, Letter
to YNP Supt.
1915
13 August
Tower Station
Record
Cruse Black
(1315-16)
Donald Stevenson
(1915-16)
Black
Stevenson
Black
Stevenson
Black
Supt . Annual
Report (1915)
1915
3 September
1915
10 October
16 October
1915
19 October
1915
23 October
24 October
28 October
30 October
1915
2 November
3 November
1915
3 November
1915
22 November
1915
6 December
1915
Stevenson
1916
6 January
Band of apparently 8 or 10 large
wolves ranging on the upper part
of Pelican Creek .... There
are also some wolves along Slough
Creek and some old and young
along Lamar River. Tnere see~
to be very few elk calves left
where these wolves range.
One wolf seen 9 miles east of
Tower .
One black wolf seen between
Tower Falls and Buffalo Ranch
(Lamar) .
Two-year-old cow elk killed by
wolves between Buffalo Ranch
(Lamar) and west Lamar Canyon.
One wolf track between Lake and
Pelican Cabin.
Trapped one female wolf (Rose
Creek area) .
One elk killed by wolves between
Buffalo Ranch (Lamar) and Black-
tail.
Killed one black female wolf up
Slough Creek.
Killed one black male wolf in
Slough Creek.
Tracks of nine wolves up
Pelican Creek from cabin.
A few wolf tracks seen in Pelican
and Raven Creeks.
One elk calf killed by wolves
between Buffalo Creek and
Specimen Ridge.
One wolf track seen 5 miles down
valley from Pelican Cabin.
One cow elk killed by gray
wolves.
Gray wolves are increasing and
have become a decided menace to
the herds of elk, deer, mountain
sheep, and antelope. Several
were killed in the Park last
winter, and an effort will be
made the coming winter to capture
or kill them.
Saw three wolves in Geode Creek
Canyon. Killed one.
Natural Resources Report
31
Appendix II
Wolf Reports 191-5-26
Source
Date
Report
Black
Tower Station
Record
Stevenson
Black
Bailey (1930)
Tower Station
Record
Stevenson
Bailey (1930)
1916
7 January
17 January
18 January
1916
18 January
1916
20 January
1916-
31 January
1916
January
1916
13 February
1916
14 February
19-20 March
22 March
26 March
1916
26 March
Tracks of three wolves between
Tower and Buffalo Ranch (Lamar) .
Wolf tracks on Specimen Ridge.
One adult cow elk killed by
wolves.
Followed tracks of four wolves
for 10 miles on Specimen Ridge
and found one adult cow elk
killed by them.
One wolf seen between Tower and
Slough Creek.
Tracks of two wolves in the
Blacktail area.
One old cow elk killed by
wolves .
In January, 1916, they [wolves]
were found in the Lamar and
Yellowstone Valleys, where
Stevenson and Black secured
four of the old wolves and,
later, a family of seven.
Two wolves seen between Tower
and lower Yellowstone River.
Tracks of two wolves seen in
Blacktail area.
Wolf tracks between Blacktail
Cabin and He 11 roaring Cabin.
Wolf tracks between Hellroaring
and Buffalo Ranch (Lamar) .
Hunted wolf dens. One wolf
tracked to den near Hellroaring
. . . female seen.
One [den] found by Stevenson and
Black on the rough slope near
Hellroaring Creek on March 26,
watched for some days in an
effort to shoot the old wolves,
which finally became suspicious
and carried the pups away to
another location farther up the
side of the mountain. The den
was described as composed of four
or five large burrows dug into
the open hillside and had
evidently been used for several
years as a score or more of old
elk skulls were lying about,
and one fresh elk head that had
recently been brought in was
found.
National Park Service
Appendix II
Wolf Reports 1914-26
source
Date
Resort
otevenson
Bailey
(1930)
Stevenson
Supt . Annual
Report (1916)
1916
27 March
28 March
30 March
14 April
15 April
16 April
1916
16 April
1916
19 April
29 April
30 April
1 May
5 May
10 May
12 May
1916
One wolf seen near Hellroaring.
Two wolves howling near Hellroaring
One large white wolf seen near
Hellroaring .
Dug out wolf den in the Hellroaring
area but they had moved.
Found wolf den.
One wolf seen and six pups caught.
On April 14 [sis], this family sf
wolves was located about s rrrile
from the first dev. in a natural
cave among some loose rocks.
Back about eight feet from the
entrance of the cave seven wolf
pups estimated to be three weeks
old were secured. A freshly killed
young elk was found about a half
mile from the den and there were
pieces of elk meat in the den
with the pups. The old wolves
were very shy and kept well out
of sight while the den was being
watched but were frequently heard
howling and answering each other
from different points snd the old
male was several times seen guard-
ing the den from a point high
above. The male is consistently
larger than the female, weighing
well over a hundred pounds.
Wolf tracks between Tower and
Blacktail .
Wolf den found near Hellroaring,
and one old wolf shot.
One pup dug out.
Tracks near Hellroaring Creek.
Tracks near Yancey's.
Den found between Slough Creek
Cabin and Hellroaring Cabin.
Den found between Yancey's and the
Buffalo Ranch (Lamar). One wolf
pup dug out.
From October 6, 1916 to June 30,
1916, two United States Biologi-
cal Survey hunters killed 12
wolves . . . skulls sent to the
National Museum. Two young male
Natural Resources Report No. 14
Appendix II
Wolf Reports 1914-26
Source
Date
Report
Supt. Annual
Report (1916) - cont.
Supt. Monthly
Report
Supt. Monthly
Report
Supt. Annual
Report (1918)
Supt. Monthly
Report
1917
December
1918
January
February
March
April
Supt. Annual
Report (1919)
May
June
July
August
September
October
November
December
1918
1919
January
February
November
1919
wolves captured in the spring
of 1915 by the employee at the
buffalo farm [Lamar] were shipped
alive on November 16 to the
National Zoological Park.
Wolves not numerous . . . two
killed by lion hunter Elkins.
Wolves reported in several
different sections of the Park.
Pack of about 16 wolves on
Specimen Ridge.
Signs of wolves on Specimen
Ridge.
Nineteen wolves killed, with
indications of many more on
Specimen Ridge and He 11 roaring.
Seven wolves killed. Towards
the end of the month the wolves
seemed to leave the Specimen
Ridge district and have not
been much in evidence since.
Four wolves killed.
One large gray wolf killed.
Sign of two wolves in the
upper Yellowstone area reported
by Biological Survey hunter
demons.
One gray wolf trapped and shot.
Three gray wolves killed.
No wolves killed but they were
considerably in evidence on
Slough and Hellroaring Creeks.
No wolves killed but sign found
along north line.
Thirty-six wolves killed in the
park the year.
No wolves killed, sign along
north line.
Two wolves killed. Signs indi-
cate presence of several ranging
from Mammoth to Soda Butte . . .
efforts are being made to
exterminate them.
Reports received of wolves
present.
Six wolves killed in the
park.
National Park Servi
Appendix II
Wolf Reports 1914-26
Source
Date
Report
Supt. Monthly
Report
Supt. Annual
Report (1920)
Supt. Monthly
Report
Supt . Annual
Report (1922)
Supt. Monthly
Report
1920
January
March
April
November
1920
1921
January
February
March
April
October
November
December
1921-22
1922
February
April
May
Three wolves, including two
females, killed.
Nine wolves killed . . . one in
the northeast and eight (including
seven pups) in Blacktail .
At least 14 wolves killed . . ..
one adult in the northeast,
eight pups in a den near Tower
Falls, five pups in another den
near Tower Falls, and one den
with pups closed up solid on
Blacktail Deer Creek.
Pack of nine wolves and tracks
seen near Tower Falls.
Twenty-eight wolves killed by
two rangers.
Forty coyotes and wolves (not
distinguished) killed.
Two black wolves seen in Slough
Creek and Specimen Ridge.
Several wolf dens located and
kept under surveillance.
One den dug out and male
(largest ever) and 11 pups
destroyed.
One wolf killed by Anderson.
One wolf killed, and many
tracks seen in Mammoth and
Blacktail Districts.
A few wolves identified from
signs.
Twenty-four wolves killed.
One wolf killed during the
winter.
Wolf dens located first week
of April between Blacktail and
Hellroaring, and adult female
killed and 10 pups captured
alive. Thirteen (sic) wolves
taken to date.
One wolf den discovered on
Specimen Ridge, destroyed both
adults and six pups. Felt that
this was the pair ranging in
Lamar Valley for several years
— 21 (sic) wolves killed to
date. It is evident that the
work of controlling these ani-
mals must be vigorously
Natural Resources Report No. 1»+
35
Appendix II
Wolf Reports 1914-26
Source
Date
Report
Supt. Monthly
Report - Cant.
July
August
October
Supt. Monthly
Report
Supt. Annual
Report (1923)
Supt. Monthly
Report
1923
April
December
1923
1924
January
February
March
prosecuted by the most effective
means available whether or not
this meets with the approval of
certain game conservationists.
Three wolves killed.
Park Ranger Henry Anderson has
been hunting out the summer
haunts of park wolves and has
succeeded in finding what he
believes to be their main sum-
mer range. This is in a section
of the park that is practically
inaccessible due to bog holes,
rim rock, down timber and jack
pines. The area is the part of
the Mirror Plateau lying near
the head of Timothy, Raven,
Pelican and Broad Creeks. Ranger
Anderson found numerous signs in
this region and due to its
inaccessibility and the fact
that there is a large amount
of game making its summer range
near this point, there is no
reason why the wolves should
not find this area ideal summer
home. Anderson will spend the
remainder of the month in this
locality in an attempt to ex-
terminate as many of these preda-
tory animals as possible.
Two large wolves, one gray and
one black, were killed by Henry
Anderson near confluence of
Pelican and Raven Creeks (these
are the two mounted specimens
in the park museum) .
Wolf den near Tower Falls cleared
out; old female killed and five
pups brought out alive to Mammoth
for exhibition.
Wolf signs have been seen near
Soda Butte and on Pelican Creek.
Eight wolves killed.
Wolf seen near Bridger Lake on
January 15 ... . Wolf signs
have been very rare this season.
One wolf seen on Elephant Back
near Lake; wolf sign near Mud
Geyser.
Wolf signs have been very scarce
throughout the entire season
National Park Service
Appendix II
Kolf Reports 1914-26
Source
Date
ReDor':
Supt. Monthly
Report - Cont.
of any
April
May
September
Supt. Monthly
Report
Supt. Monthly
Report
Bailey (1930)
November
December
1925
October
1926
January
October
December
1926
and we have no rev or
wolf kills in the park
Signs of wolf exceedingly
saarae ....
First wolf sign reported during
past year observed near Soda
Butte . . . also at about the
same time at Pelican Creek.
There were no reported instances
of wolf activity in the park
last winter apart from an occa-
sional lone track. None were
actually seen in the park. The
situation this fall gives promise
of a recurrence of the wolf as
we have two reports of recent
date indicating their presence
in the Park. Park Ranger Hall
reports having seen three near
Heart Lake and a wolf pack num-
bering twelve are reported to
have been seen at Elk Park by
two members of a road crew on
duty near that point. This last
report has not been definitely
confirmed and may be exaggerated.
A number of wolf signs were ob-
served in October but only one
track has been reported for
November .
Two wolf signs seen on December
16 on south slope of Saddle
Mountain.
Three wolf signs were observed
on the east shore of Yellow-
stone Lake on the 20th.
One wolf sign recently seen on
Cabin Creek.
There is believed to be a very
limited number of wolves in the
Park.
Sign of one, perhaps two. wolves
along lower trail between Hell-
roaring and Tower Falls.
Tracks on Two Ocean Pass and one
wolf seen on Trident Plateau by
Sierra Club party.
Natural Resources Report No. 1U
37
APPENDIX III
Summary of Wolf Reports, 1927-36, Yellowstone National Park.
Source
Date
Report
Supt. Monthly
Report
Supt. Monthly
Report
Supt. Monthly
Report
Supt. Monthly
Report
Arnold
(1957) •
Supt. Monthly
Report
1927
February
October
November
1928
February
October
1929
September
1930
January
1934
winter
1936
April
Wolf signs have been rarely seer
in the Park this winter and we
have every reason . to consider
that there are only a very few
in the entire park area.
It is doubtful if there are more
than a very few wolves in the
entire park area.
Ranger Ogston reports signs of
two wolves in the vicinity of
the Slough Creek mailbox.
Wolf signs have been rarely
observed.
There have been no wolf sign
reported this season.
Wood crews at Yellowstone Lake
reported seeing two gray wolves.
A wolf has been reported working
in the Tower Falls and Hellroaring
districts. The tracks have been
seen many times, and two elk
calves have been found that were
from all evidence, killed by
this wolf.
Four wolves seen up Tower Creek.
Five wolves were seen and reported
in the vicinity of Old Faithful.
Several freshly killed elk car-
casses found in this vicinity
suggest that the animals seen
were actually wolves instead of
coyotes.
National Park Service
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